喀纳斯泰加林群落火成演替中草本层与林冠层物种的连锁关系

doi:10.11707/j.1001-7488.20210401

摘要/Abstract

摘要：

目的: 检验喀纳斯泰加林群落火成演替中草本层与林冠层物种的连锁关系，并探索其形成机制。方法: 以新疆喀纳斯国家级自然保护区内未受人为干扰的可识别的泰加林火成演替群落为对象，基于369个森林样地及其历史火干扰时间和火烈度调查，以环境、火干扰和林分因子为解释变量，草本层和林冠层物种多样性指数为响应变量，分别进行典范对应分析（CCA），并对草本层和林冠层的CCA排序第一轴得分值进行相关分析，探讨不同烈度火干扰后草本层与林冠层物种的连锁关系和随演替阶段的变化及其原因。结果: 1）火成演替群落的草本层与林冠层CCA排序第一轴得分值之间的相关性，在弱烈度和中烈度火干扰后达到显著水平（P=0.017 < 0.05，P=0.043 < 0.05），在强烈度火干扰后未达到显著水平（P=0.093>0.05）；2）在强烈度火干扰后，火成演替群落的草本层与林冠层CCA排序第一轴得分值之间的相关性在演替前期达到显著水平（P=0.044 < 0.05），在演替中期和后期达到极显著水平（P=0.008 < 0.01，P=0.006 < 0.01）；中烈度火干扰后，相应得分值的相关性在演替前期达到显著水平（P=0.043 < 0.05），在演替中期和后期达到极显著水平（P=0.006 < 0.01，P=0.004 < 0.01）；弱烈度火干扰后，相应得分值的相关性在不同演替阶段均达到极显著水平（P=0.006 < 0.01，P=0.005 < 0.01，P=0.002 < 0.01）；3）草本层和林冠层具有一致性响应且达到显著水平的因子，在强烈度火干扰后的演替前期、中期和后期分别有1、2和4个，在中烈度和弱烈度则均为3、4和5个。结论: 火干扰后，泰加林群落演替过程中的草本层与林冠层物种之间存在连锁关系，其原因可能是草本层和林冠层物种多样性格局对相同环境梯度的响应具有一致性。该连锁关系随火烈度增强而从强到弱，且随演替向前发展变得更加紧密。

关键词: 泰加林, 火干扰, 林冠层, 草本层, 物种, 连锁关系

Abstract:

Objective: The aim of this study is to test the linkage between herbaceous layer and overstorey species along the pyrogenic successions in Kanas taiga communities and to reveal its the formation mechanism. Method: Community characteristics, i.e., fire time, fire severity, stand factors and environmental factors, of 369 samples of the identifiable unhuman-disturbed pyrogenic succession communities were investigated in Kanas taiga, Xinjiang, China. Then, the canonical correspondence analysis (CCA) and correlation analysis were performed to analyze the linkage between herbaceous layer and overstorey species. Result: The correlation in the first axis score of CCA between herbaceous layer and overstorey reached a significant level after low-severity fire and moderate-severity fires (P=0.017 < 0.05, P=0.043 < 0.05), but it did not reach a significant level after high-severity fire (P=0.093>0.05). After high-severity fire, the correlation in the first axis score of CCA between herbaceous layer and overstorey reached a significant level in early succession stage (P=0.044 < 0.05), whereas reached a highly significant level in the middle and late succession stages (P=0.008 < 0.01, P=0.006 < 0.01). After moderate-severity fire, the correlation of the first axis score of CCA between herbaceous layer and overstorey reached a significant level in early succession stage (P=0.043 < 0.05), whereas reached a highly significant level in the middle and late succession stages (P=0.006 < 0.01, P=0.004 < 0.01). After the interference of low-intensity fire, there was the highly significant correlation in the first axis score of CCA between herbaceous layer and overstorey across all succession stages (P=0.006 < 0.01, P=0.005 < 0.01, P=0.002 < 0.01). The results of CCA showed that the number of same explanatory variables between herbaceous layer and overstorey (i.e., fire time, fire severity, stand factors and environmental factors) with the significant influences on response variables (i.e., species diversity of herbaceous and canopy layers) were 1, 2 and 4 in early, middle and late succession stages after high-severity fire, respectively. However, after moderate- and low-severity fire, the number of same explanatory variables was 3, 4 and 5 in early, middle and late succession stages, respectively. Conclusion: Our results have demonstrated that after fire disturbance there is a linkage between herbaceous layer and overstorey species during community succession in Taijia forest. The reason may be that species diversity patterns in herbaceous layer and canopy layer have consistent response to same environmental gradient across forest succession. The linkage in species diversity between herbaceous layer and overstorey decreases with the intensity of fire severity, whereas increases along forest succession.

Key words: taiga, fire disturbance, overstorey, herbaceous layer, species, linkage

中图分类号:

S718.54

郭珂,潘存德,余戈壁,李贵华,张帆,邹卓颖,刘博. 喀纳斯泰加林群落火成演替中草本层与林冠层物种的连锁关系[J]. 林业科学, 2021, 57(4): 1-13.

Ke Guo,Cunde Pan,Gebi Yu,Guihua Li,Fan Zhang,Zhuoying Zou,Bo Liu. Linkage between Herbaceous Layer and Overstorey Species along the Pyrogenic Successions in Kanas Taiga Communities[J]. Scientia Silvae Sinicae, 2021, 57(4): 1-13.

图/表 11

图1

图2

图3

表1

强烈度火干扰后泰加林群落不同演替阶段草本层与林冠层物种CCA排序轴相关系数①"

因子 Factors	演替前期 Early stage of succession				演替中期 Middle stage of succession				演替后期 Late stage of succession
	草本层 Herbaceous layer		林冠层 Overstorey		草本层 Herbaceous layer		林冠层 Overstorey		草本层 Herbaceous layer		林冠层 Overstorey
	Ax1	Ax2	Ax1	Ax2	Ax1	Ax2	Ax1	Ax2	Ax1	Ax2	Ax1	Ax2
Elev	-0.360^*	0.285	-0.110	0.051	0.361^*	0.113	-0.163	0.167	0.275	-0.064	-0.041	-0.111
Slop	0.166	-0.171	0.295	0.066	0.203	0.048	-0.147	0.063	-0.131	-0.090	-0.047	0.030
TRASP	-0.069	-0.233	-0.038	0.019	0.003	-0.197	-0.085	0.031	0.013	0.004	-0.067	0.053
PFT	0.011	0.213	-0.048	-0.001	-0.175	-0.038	0.082	-0.291	-0.086	0.102	-0.085	-0.186
pH	0.264	-0.395^*	0.074	-0.051	-0.376^*	-0.207	0.050	-0.109	-0.430^**	0.118	0.189	-0.047
Con	-0.190	0.033	0.151	-0.020	0.077	-0.068	0.117	-0.158	-0.197	0.142	0.271	-0.040
SBD	0.220	0.120	-0.279	0.286	-0.136	0.310^*	-0.079	0.030	-0.100	-0.188	-0.125	-0.157
Poro	-0.334^*	0.052	0.268	-0.040	0.213	-0.310^*	0.205	-0.040	0.246	0.020	-0.211	-0.022
Org	0.009	-0.452^**	0.264	0.054	-0.261	-0.031	-0.147	0.113	-0.071	-0.221	0.099	0.155
TN	0.110	-0.244	0.328^*	0.125	-0.141	-0.042	0.363^*	0.174	-0.352^*	-0.088	0.350^*	0.087
TK	0.163	-0.201	0.068	0.196	-0.140	-0.305^*	0.193	0.160	-0.027	-0.081	-0.066	0.097
TP	0.267	-0.375^*	0.070	0.065	-0.259	-0.150	-0.222	0.048	-0.359^*	0.137	-0.359^*	0.088
AN	0.014	-0.238	0.154	-0.010	-0.081	0.012	0.130	0.131	-0.181	-0.089	0.330^*	0.103
AK	0.145	0.083	-0.320^*	0.033	-0.047	0.312^*	-0.312^*	0.146	-0.018	-0.308^*	0.100	0.138
AP	0.181	0.118	-0.421^**	0.073	0.026	0.149	-0.311^*	0.184	0.116	-0.087	0.113	0.168
Fe	-0.176	-0.154	0.066	0.143	0.118	-0.105	-0.093	0.061	0.117	-0.313^*	-0.376^*	0.188
Cu	-0.091	-0.345^*	0.483^**	-0.322^*	-0.316^*	-0.220	-0.048	-0.521^**	-0.236	0.324^*	0.014	-0.166
Zn	0.111	-0.119	0.062	0.054	-0.204	0.309^*	-0.181	0.091	-0.167	-0.101	0.316^*	0.165
Mn	0.086	0.002	-0.031	0.410^*	0.094	0.113	0.225	0.443^**	0.110	-0.013	0.183	0.071
Mg	-0.348^*	0.309	-0.067	0.188	0.211	0.086	0.145	0.154	0.232	-0.212	0.085	-0.150
Ca	-0.130	0.473^*	-0.110	0.155	0.327^*	0.149	0.088	0.016	0.273	-0.153	0.146	-0.227
LAI	0.207	-0.065	0.248	0.304	0.199	-0.065	-0.150	-0.051	0.016	0.221	-0.080	0.321^*
P_c	0.163	-0.037	0.160	0.185	0.230	-0.114	-0.157	-0.068	0.012	0.307^*	-0.108	0.317^*
DBH	-0.129	-0.145	-0.325^*	0.036	-0.051	-0.142	-0.017	0.143	0.078	-0.042	0.066	0.005
D_t	0.420^*	0.375^*	-0.025	0.072	0.146	0.149	-0.064	-0.227	-0.168	-0.192	-0.228	-0.015
H_D	-0.045	0.099	-0.452^**	0.089	0.118	0.036	-0.052	0.196	0.243	-0.132	0.159	0.053

表1

表2

中烈度火干扰后泰加林群落不同演替阶段草本层与林冠层物种CCA排序轴相关系数"

因子 Factors	演替前期 Early stage of succession				演替中期 Middle stage of succession				演替后期 Late stage of succession
	草本层 Herbaceous layer		林冠层 Overstorey		草本层 Herbaceous layer		林冠层 Overstorey		草本层 Herbaceous layer		林冠层 Overstorey
	Ax1	Ax2	Ax1	Ax2	Ax1	Ax2	Ax1	Ax2	Ax1	Ax2	Ax1	Ax2
Elev	-0.515^**	-0.033	-0.056	-0.306^*	0.137	0.336^*	-0.345^*	0.056	-0.458^**	0.044	-0.346^*	-0.098
Slop	0.204	0.005	0.115	-0.067	0.106	-0.011	0.235	0.025	-0.107	0.064	-0.084	0.008
TRASP	-0.034	0.054	0.045	-0.019	0.153	-0.101	0.107	0.165	-0.172	-0.046	0.033	-0.099
PFT	-0.089	-0.006	0.241	-0.059	-0.043	0.007	-0.283	0.159	-0.087	0.109	0.067	0.059
pH	0.336^*	-0.072	0.062	0.281	0.057	0.048	0.214	-0.036	0.142	-0.015	0.134	0.097
Con	0.322^*	0.045	0.083	0.206	-0.096	-0.266	-0.123	0.161	-0.318^*	-0.112	0.103	0.060
SBD	0.180	-0.117	-0.125	0.136	0.026	0.297	-0.024	-0.069	0.141	0.219	-0.253	-0.153
Poro	-0.547^**	0.111	-0.061	-0.345^*	-0.227	0.133	-0.612^**	-0.175	-0.206	0.015	0.038	0.189
Org	0.067	0.373^*	0.303	-0.125	-0.125	-0.250	-0.174	0.437^**	-0.165	0.021	0.361^*	-0.131
TN	0.003	0.439^**	0.325^*	-0.177	-0.330^*	-0.196	-0.484^**	0.379	-0.407^*	-0.102	0.407^*	-0.256
TK	-0.413^*	0.005	-0.141	-0.210	-0.131	-0.358^*	-0.275	0.062	-0.324^*	0.063	0.203	-0.383^*
TP	-0.006	0.230	0.120	-0.147	0.020	0.195	-0.154	0.089	0.164	-0.342^*	0.378^*	-0.141
AN	0.105	0.427^**	0.202	-0.073	-0.073	-0.311^*	0.015	0.306	-0.052	-0.163	0.381^*	-0.091
AK	0.020	0.259	0.082	0.102	-0.152	0.066	0.286	0.043	-0.038	-0.011	0.156	-0.046
AP	-0.174	-0.042	-0.084	0.099	0.337^*	0.247	0.643^**	0.080	0.085	0.117	-0.372^*	-0.238
Fe	-0.180	0.058	-0.334^*	-0.063	0.035	0.149	0.492^**	-0.205	-0.246	0.387^*	-0.174	-0.348^*
Cu	0.420^**	0.109	0.200	-0.108	0.057	-0.025	-0.269	0.125	-0.117	-0.450^*	0.001	0.068
Zn	0.203	0.245	0.326^*	0.104	-0.251	0.047	-0.001	0.228	-0.087	-0.108	-0.069	-0.386^*
Mn	-0.124	0.011	0.156	-0.007	-0.274	0.013	-0.165	0.196	-0.192	0.000	0.058	-0.196
Mg	-0.434^**	0.163	-0.165	-0.159	-0.353^*	-0.275	-0.370^*	-0.228	-0.280	0.004	0.072	0.174
Ca	-0.107	-0.133	-0.048	0.333^*	-0.155	-0.340^*	-0.094	-0.037	-0.224	0.054	0.106	0.064
LAI	0.443^**	-0.241	-0.082	0.077	-0.231	0.051	-0.120	-0.081	0.240	-0.036	-0.111	0.135
P_c	0.416^**	-0.221	-0.025	0.063	-0.250	0.078	-0.091	-0.125	0.166	0.122	-0.140	0.184
DBH	0.234	-0.020	0.005	-0.025	-0.071	-0.052	-0.324^*	0.294	0.096	-0.159	0.212	-0.079
D_t	-0.076	-0.009	0.065	-0.197	0.214	-0.113	-0.022	-0.024	0.094	-0.001	-0.175	-0.127
H_D	0.181	-0.026	-0.085	0.156	-0.225	0.065	-0.357^*	0.094	0.029	0.175	-0.229	-0.052

表2

表3

弱烈度火干扰后泰加林群落不同演替阶段草本层与林冠层物种CCA排序轴相关系数"

因子 Factors	演替前期 Early stage of succession				演替中期 Middle stage of succession				演替后期 Late stage of succession
	草本层 Herbaceous layer		林冠层 Overstorey		草本层 Herbaceous layer		林冠层 Overstorey		草本层 Herbaceous layer		林冠层 Overstorey
	Ax1	Ax2	Ax1	Ax2	Ax1	Ax2	Ax1	Ax2	Ax1	Ax2	Ax1	Ax2
Elev	-0.578^**	0.318^*	-0.197	0.341^*	-0.333^*	0.268	-0.239	-0.423^**	-0.342^*	-0.107	-0.385^*	0.126
Slop	-0.260	-0.089	0.152	0.386^*	-0.033	0.125	0.044	0.146	0.103	-0.217	0.001	-0.056
TRASP	0.182	0.119	0.352^*	0.155	0.127	0.199	0.007	-0.122	0.060	-0.073	0.131	-0.043
PFT	0.087	0.324^*	0.364^*	0.239	0.244	0.268	-0.126	-0.201	0.340^*	-0.025	-0.019	-0.187
pH	0.088	0.228	-0.181	-0.025	0.129	-0.270	-0.123	0.217	-0.110	0.021	-0.037	-0.153
Con	-0.071	-0.008	-0.341^*	-0.135	-0.076	0.044	-0.307^*	0.140	-0.016	-0.078	0.185	-0.139
SBD	-0.037	-0.021	0.161	0.112	0.029	0.463^**	0.074	0.040	-0.017	-0.200	-0.046	0.094
Poro	-0.200	0.114	0.191	0.077	-0.380^*	0.342^*	-0.134	-0.086	0.019	0.303^*	-0.057	-0.143
Org	-0.093	-0.055	-0.009	-0.187	0.160	-0.530^**	-0.323^*	0.260	-0.020	-0.149	0.304^*	-0.107
TN	-0.254	-0.077	0.303	-0.254	-0.109	-0.016	-0.202	0.190	-0.018	-0.107	0.368^*	0.021
TK	-0.350^*	-0.193	0.011	0.208	-0.364^*	0.160	-0.086	-0.050	-0.020	0.302^*	-0.169	0.324^*
TP	-0.257	-0.152	0.079	0.078	0.207	-0.119	-0.163	0.119	0.114	-0.264	-0.014	-0.116
AN	0.077	-0.318^*	0.167	-0.070	0.090	0.009	-0.221	0.320^*	-0.020	-0.004	-0.020	-0.023
AK	-0.257	0.069	-0.451^**	0.028	0.099	0.178	0.159	0.210	-0.014	0.201	-0.065	0.076
AP	0.164	0.237	0.113	0.023	0.281	0.120	0.104	0.315	0.022	0.301^*	-0.309^*	-0.100
Fe	0.125	-0.188	0.176	-0.390^*	0.091	-0.003	0.057	0.106	0.002	-0.155	0.021	0.059
Cu	0.089	-0.060	-0.035	-0.273	-0.134	-0.624^**	-0.320^*	-0.121	-0.091	-0.534^**	0.358^*	-0.198
Zn	-0.414^*	-0.345^*	-0.145	0.008	-0.065	-0.458^**	0.040	0.103	-0.125	-0.345^*	0.034	0.074
Mn	-0.176	0.184	0.037	0.111	-0.082	0.270	0.040	0.093	0.042	-0.135	-0.003	-0.136
Mg	-0.054	0.166	-0.041	0.176	-0.427^**	0.331^*	-0.058	-0.079	0.111	-0.164	-0.015	0.131
Ca	0.226	0.437^**	-0.055	-0.064	-0.100	0.170	-0.005	-0.064	-0.091	-0.058	0.133	0.063
LAI	0.187	-0.140	-0.074	0.177	0.141	0.285	-0.101	0.086	0.129	0.004	0.136	-0.115
P_c	0.239	-0.114	-0.097	0.195	0.158	0.234	-0.096	0.107	0.160	0.010	0.188	-0.124
DBH	-0.180	0.059	-0.274	-0.262	-0.017	-0.041	-0.184	-0.140	0.082	-0.260	0.083	-0.138
D_t	0.462^**	-0.086	0.490^**	-0.002	-0.433^**	-0.185	-0.317^*	-0.108	0.066	-0.174	0.123	-0.133
H_D	0.028	0.024	-0.544^**	-0.103	0.009	-0.047	-0.321^*	0.275	0.341^*	-0.180	-0.101	-0.311^*

表3

图4

图5

图6

图7

图8

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